Darwinism in Forestry

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DARWINISM IN FORESTRY

RAPHAEL ZON

U. S. Forest Service

The centennial anniversary of the birth of Charles
Darwin was the occasion for many interesting reviews of
what Darwinism has clone for the biological sciences. In
all these reviews, however, scarcely any reference is made
to forestry. Yet historically and inherently there is a
most remarkable and unique connection between Darwin-
ism and forestry.

On April 10, 1860, soon after the appearance of the first
edition of the "Origin of Species," Darwin wrote to his
friend C. Lyell :

Now for a curious thing' about my book, and then I have done. In
last Saturday's Gardeners' Chronicle, a Mr. Patrick Matthew publishes
a long extract from his work on " Naval Timber and Arboriculture,"
published in 1S31, in which he briefly but completely anticipates the
theory of Natural Selection. I have ordered the book, as some few
passag-es are rather obscure, but it is certainly, I think, a complete but
not developed anticipation ! One may be excused in not having dis-
covered the fact in a work on Naval Timber. 1

And three days later, on April 13, 1860, he wrote to J.
D. Hooker. 3

My dear Hooker — Questions of priority so often lead to odious
quarrels, that I should esteem it a great favor if yon would read the
enclosed. If yon think it proper that I should send it (and of this
there can hardly be any question), and if you think it full and ample
enough, please alter the date to the day on which you post it, and let-
that be soon. The case in the Gardeners' Chronicle seems a little
stronger than in Mr. Matthew's book, for the passages are therein
scattered in three places; but it would be mere hair-splitting to notice
that. If you object to my letter, please return it; but I do not expect
that you will, but I thought that you would not object to run your eye
over it.

'"The Life and Letters of Charles Darwin," by F. Darwin, 1898, New
York, Appleton & Co., p. 95.
■ll'ul., pp. 95 and 96.

540

No. 561] DARWINISM IN FORESTRY 541

The statement to which Darwin referred in his letter
to Hooker appeared in the Gardeners' Chronicle on April
21, 1860 (page 362) , and is this :

I have been much interested by Mr. Patrick Matthew's communica-
tion in the number of your paper dated April 7th. I freely acknowl-
edge that Mr. Matthew has anticipated by many years the explanation
which I have offered of the origin of species, under the name of natural
selection. I think that no one will feel surprised that neither I, nor
apparently any other naturalist, had heard of Mr. Matthew's views,
considering how briefly they are given, and that they appeared in the
appendix to a work on Naval Timber and Arboriculture. I can do no
more than offer my apologies to Mr. Matthew for my entire ignorance
of this publication. If another edition of my work is called for, I will
insert to the foregoing effect. 3

In the Historical Sketch 4 which he added to the later
editions of his book Darwin gives Matthew credit for the
Nature's law of selection in the following words:

In 1831 Mr. Patrick Matthew published his work on " Naval Timber
and Arboriculture," in which he gives precisely the same view on the
origin of species as that (presently to be alluded to) propounded by
Mr. Wallace and myself in the Linnean Journal, and as that enlarged
in the present volume. Unfortunately, the view was given by Mr.
Matthew very briefly in scattered passages in an Appendix to a work
on a different subject, so that it remained unnoticed until Mr. Matthew
himself drew attention to it in the Gardeners' Chronicle, on April 7th,
1860. The differences of Mr. Matthew's view from mine are not of
much importance: he seems to consider that the world was nearly
depopulated at successive periods, and then re-stocked ; and he gives as
an alternative, that new forms may be generated " without the presence
of any mould or germ of former aggregates." I am not sure that I
understand some passages; but it seems that he attributes much in-
fluence to the direct action of the conditions of life. He clearly saw,
however, the full force of the 'principle of natural selection. 3

In a letter written by Darwin to J. L. A. de Quatref ages
on April 25, 1861, he referred to Patrick Matthew's ex-
planation in a postscript as follows :

I have lately read M. Naudin's paper, but it does not seem to me to
anticipate me, as he does not show how selection could be applied under

3 Ibid.

""The Origin of Species," 1878, p. xvi— Historical Sketch.

5 Ibid.

512 THE AMERICAN NATURALIST [Vol. XLVII

nature ; but an obscure writer on forest trees, in 1830, in Scotland, most
expressly and clearly anticipated my views — though he put the case so
briefly that no single person ever noticed the scattered passages in his
book.

Grant Allen in his biography of Darwin (1888) calls
Patrick Matthew the unconscious author of the principle
of natural selection which he applied in his book on naval
timber to the entire Nature.

Here then is a most interesting fact which seems to me
of deep significance to foresters. The first Darwinian,
who twenty-nine years before Darwin formulated the law
of natural selection, was a forester. I shall not attempt
here to compare Darwin's and Matthew's views on nat-
ural selection. Matthew's book, the full title of which is
"Naval Timber and Arboriculture, With Critical Notes
on Authors Who Have Eecently Treated the Subject of
Planting," is accessible in the Congressional Library.
The chapter on Nature's Law of Selection I hope can be
reprinted in the next issue of the Proceedings of the So-
ciety of American Foresters, so that every one will be
able to draw the comparison for himself.

In bringing together this evidence I am very far indeed
from any desire to detract in the least from the great
service which Darwin rendered to science. It was Dar-
win who first gave flesh and blood to the idea of natural
selection. It was his wonderful interpretation of all bio-
logical facts in the light of natural selection that made the
latter the universal law applicable to the entire organic
world. Before this accomplishment the claims of all
others must sink into obscurity.

My purpose in assembling these records is twofold:
First, to restore the memory of one who ploughed the
same fields as we do now, the name of a forester whose
idea, although it did not perish, slumbered almost un-
known for nearly thirty years until another and bigger
man brought it to life and general recognition; and sec-
ond, to offer an explanation of the reason why a forester
above all others should be the one to observe and formu-

No. 561] DARWINISM IN FORESTRY 543

late the law of the struggle for existence as the basis for
natural selection and the origin of new species.

My first purpose, I hope, has been accomplished by
quoting extracts from Darwin's correspondence. The
second still remains.

There is nothing accidental, in my opinion, in the fact
that a forester should be the first to observe the struggle
for existence and its bearing upon the development of the
new varieties, because there is no other plant society in
the world which presents a more striking example of the
struggle for existence and of natural selection than the
forest. Nowhere else, also, can the law of this process be
more fully studied.

The regular decrease in the number of trees on a given
area with increase in age forms one of the earliest obser-
vations of the foresters, who, at a time antedating Dar-
win, properly gave this process the name of the struggle
for existence, the struggle for the necessary growing
space. The foresters have discovered the laws governing
this process, a process in which almost 95 per cent, of all
trees that start life in the stand perish, and in the form
of yield tables have expressed it quantitatively, have
measured and weighed it. They have shown how this
struggle for existence varies with the species, climate,
drainage and soil conditions, and age of the stand ; that it
is more intense, and consequently the differentiation into
dominant and suppressed classes occurs earlier with
light-needing species than with shade-enduring ones. In
a climate most suitable to the species and on favorable
situations this struggle again results in more rapid dif-
ferentiation into dominant and suppressed trees than
when the species grow outside of their optimum range
and on poor soils. These are elementary and fundamen-
tal facts known to foresters for many years.

The foresters have not only observed these facts, but
they have also furnished an explanation for them. The
more favorable the conditions of growth, the greater is
the development of the individual trees; the earlier,

544 THE AMERICAN NATURALIST [Vol. XLYII

therefore, begins the struggle for space and the differen-
tiation into dominant and suppressed, with the subse-
quent dying out of the latter. They have followed this
process throughout the entire life of the stand, have es-
tablished its various degrees of severity, and have dis-
covered its culmination during the period of the most
rapid growth in height. This struggle for space and
light is the basis of the forester's operations, as only by
utilizing and controlling it is he capable of producing
wood of high technical qualities, tall cylindrical boles,
free of branches, and wood with uniform annual rings
possessing great elasticity. Without this struggle there
is no forest, there is no production of valuable timber,
save firewood.

The struggle for existence in a forest stand is not con-
fined to individual members of the same age or the same
story, but the forest, as a whole, battles for its existence
against the adjoining meadow, swamp or shrub vegeta-
tion ; the old trees against the young growth that comes
up under them ; groups of trees of different species or of
different ages against each other. In this struggle the
forest accomplishes what no other vegetation does ;
namely, it actually changes the climate over the area oc-
cupied by it, and makes it inhospitable for its enemies.
The forest creates its own interior environment to which
its own members are completely adapted, but in which
other species find either too much or too little light, the
humus too scant or too deep, or too acid, the temperature
too high or too low. Whatever it may be, the forest's
competitors are eliminated through the changed environ-
ment. To change this environment, however, there must
be a close stand, there must be present the struggle for
existence among the individual members of the stand.
Through interior struggle among its own members the
stand secures resistance against invasion by other vege-
tation. How manifoldly broad and deep, then, is the
struggle for existence in the forest.

When we come now to natural selection nowhere else is

No. 561] DARWINISM IN FORESTRY 545

it expressed in such fullness and so strikingly as in the
forest. The forest is a natural breeding place in which
constantly only the trees best adapted to the climate and
the situation are allowed to remain. In the forest only
the conquerors in the struggle for existence are the ones
which produce seed in abundance. During a seed year
the dominant and co-dominant trees produce seed in large
quantities ; the intermediate trees, which may properly
be called the candidates for suppression, participate but
little, and then only in exceptionally good seed years,
while the oppressed and suppressed do not bear seed at
all. With what rigidity, then, must the natural selection
go on in a forest, if we consider first what a small per-
centage of trees in a stand of the same generation come to
be conquerors in the struggle for existence; second, the
great age reached by trees ; third, the numerous genera-
tions of trees that have succeeded each other in the same
forest ; and fourth, the relatively limited capacity of tree
seeds for dissemination. With each generation the for-
est trees must become more and more delicately adjusted
and adapted to the given conditions of growth. The new
generation inevitably arises from seed sown by the best
developed trees, from those which have withstood the long
and intense battle not only against Nature alone, but
against Nature in the presence of competitors. Of this
possibly only 1 per cent, or less will reach maturity and
be able to continue the species. No wonder, therefore,
that in spite of search for new species all over the world
so few forest trees have been successfully introduced into
new countries and so little progress has been made with
the artificial improvement of them. So perfect is the nat-
ural selection in the forest, so fine is the adjustment be-
tween the environment and the forest trees, that it is al-
most impossible for man to approach it. I do not mean
the introduction of trees for park purposes or breeding
new varieties for some other purpose than timber; I have
in mind only the establishment of natural forests and the
production of timber.

546 THE AMEBIC AN NATURALIST [Vol. XL VII

The natural selection forms also the basis of the for-
ester's operation in selecting trees for seeding purposes,
in making regeneration cuttings, in collecting seed for re-
forestation and so on.

These few facts are enough to show with what fullness
and force the principles advanced by Darwin are ex-
pressed in the forest. If agriculture furnished Darwin
with many examples of artificial selection upon which he
built by analogy his principle of natural selection, the
forest, of all plant formations, furnishes the most strik-
ing examples and proof of the latter. As a matter of fact,
forestry as an art is nothing else but the controlling and
regulating of the struggle for existence for the practical
ends of man; forestry as a science is nothing else but the
study of the laws tvhich govern the struggle for existence.

Is there anything strange, therefore, that it was a for-
ester who first formulated the principles of natural se-
lection? Is there anything strange, also, in the fact that
it was also foresters who have laid the foundation for
what has come to be known as ecology, which is the log-
ical development of Darwinism? Because of the fact
that the forest is the highest expression of plant life, the
foresters occupy the strategic position from which they
command vistas accessible only with difficulty to other
naturalists. In this lies the strength of forestry, its pe-
culiar beauty, and the debt which science owes to it.